James E. O'Hara

Signal through the noise? Phylogeny of the Tachinidae (Diptera) as inferred from morphological evidence

The oestroid family Tachinidae represents one of the most diverse lineages of insect parasitoids. Despite their broad distribution, diversity and important role as biological control agents, the phylogeny of this family remains poorly known. Here, we review the history of tachinid systematics and present the first quantitative phylogenetic analysis of the family based on morphological data. Cladistic analyses were conducted using 135 morphological characters from 492 species belonging to 180 tachinid genera, including the four currently recognized subfamilies (Dexiinae, Exoristinae, Phasiinae, Tachininae) and all major tribes. We used characters of eggs, first‐instar larvae and adults of both sexes. We examined the effects of implied weighting by reanalysing the data with varying concavity factors. Our analysis generally supports the subfamily groupings Dexiinae + Phasiinae and Tachininae + Exoristinae, with only the Exoristinae and the Phasiinae reconstructed as monophyletic assemblages under a wide range of weighting schemes. Under these conditions, the Dexiinae, which were previously considered a well‐established monophyletic assemblage, are reconstructed as being paraphyletic with respect to the Phasiinae. The Tachininae are reconstructed as a paraphyletic grade from which the monophyletic Exoristinae arose. The Exoristinae are reconstructed as a monophyletic lineage, but phylogenetic relationships within the subfamily are largely unresolved. We further explored the evolution of oviposition strategy and found that the oviparous groups are nested within ovolarviparous assemblages, suggesting that ovipary may have evolved several times independently from ovolarviparous ancestors. This counterintuitive pattern is a novel hypothesis suggested by the results of this analysis. Finally, two major patterns emerge when considering host associations across our phylogeny under equal weights: (i) although more than 60% of tachinids are parasitoids of Lepidoptera larvae, none of the basal clades is unambiguously associated with Lepidoptera as a primitive condition, suggesting that tachinids were slow to colonize these hosts, but then radiated extensively on them; and (ii) there is general agreement between host use and monophyly of the major lineages.

Annotated catalogue of the Tachinidae (Insecta, Diptera) of the Afrotropical Region, with the description of seven new genera

Abstract The Tachinidae of the Afrotropical Region are catalogued and seven genera and eight species are newly described. There are 237 genera and 1126 species recognized, of which 101 genera and 1043 species are endemic to the region. The catalogue is based on examination of the primary literature comprising about 525 references as well as numerous name-bearing types and other specimens housed in collections. Taxa are arranged hierarchically and alphabetically under the categories of subfamily, tribe, genus, subgenus (where recognized), species, and rarely subspecies. Nomenclatural information is provided for all genus-group and species-group names, including lists of synonyms (mostly restricted to Afrotropical taxa) and name-bearing type data. Species distributions are recorded by country within the Afrotropical Region and by larger geographical divisions outside the region. Additional information is given in the form of notes, numbering about 300 in the catalogue section. Seven genera and eight species are described as new: Afrophylax Cerretti & O’Hara with type species Sturmia aureiventris Villeneuve, 1910, gen. n. (Exoristinae, Eryciini); Austrosolieria Cerretti & O’Hara with type species Austrosolieria londti Cerretti & O’Hara, gen. n. and sp. n. (South Africa) and Austrosolieria freidbergi Cerretti & O’Hara, sp. n. (Malawi) (Tachininae, Leskiini); Carceliathrix Cerretti & O’Hara with type species Phorocera crassipalpis Villeneuve, 1938, gen. n. (Exoristinae, Eryciini); Filistea Cerretti & O’Hara with type species Viviania aureofasciata Curran, 1927, gen. n. and Filistea verbekei Cerretti & O’Hara, sp. n. (Cameroon, D.R. Congo, Uganda) (Exoristinae, Blondeliini); Mesnilotrix Cerretti & O’Hara with type species Dexiotrix empiformis Mesnil, 1976, gen. n. (Dexiinae, Dexiini); Myxophryxe Cerretti & O’Hara with type species Phorocera longirostris Villeneuve, 1938, gen. n., Myxophryxe murina Cerretti & O’Hara, sp. n. (South Africa), Myxophryxe regalis Cerretti & O’Hara, sp. n. (South Africa), and Myxophryxe satanas Cerretti & O’Hara, sp. n. (South Africa) (Exoristinae, Goniini); and Stiremania Cerretti & O’Hara with type species Stiremania karoo Cerretti & O’Hara, gen. n. and sp. n. (South Africa), and Stiremania robusta Cerretti & O’Hara, sp. n. (South Africa) (Exoristinae, Goniini). Paraclara Bezzi, 1908 is transferred from the Cylindromyiini to the Hermyini, comb. n. Sarrorhina Villeneuve, 1936 is transferred from the Minthoini to the Graphogastrini, comb. n. Three genera are newly recorded from the Afrotropical Region: Madremyia Townsend, 1916 (Eryciini); Paratrixa Brauer & Bergenstamm, 1891 (Blondeliini); and Simoma Aldrich, 1926 (Goniini). Three genera previously recorded from the Afrotropical Region are no longer recognized from the region: Calozenillia Townsend, 1927 (Palaearctic, Oriental and Australasian regions); Eurysthaea Robineau-Desvoidy, 1863 (Palaearctic, Oriental and Australasian regions); and Trixa Meigen, 1824 (Palaearctic and Oriental regions). Two species are newly recorded from the Afrotropical Region: Amnonia carmelitana Kugler, 1971 (Ethiopia, Kenya); and Simoma grahami Aldrich, 1926 (Namibia). Three species previously recorded from the Afrotropical Region are no longer recognized from the region: Euthera peringueyi Bezzi, 1925 (Oriental Region); Hamaxia incongrua Walker, 1860 (Palaearctic, Oriental and Australasian regions); Leucostoma tetraptera (Meigen, 1824) (Palaearctic Region). New replacement names are proposed for five preoccupied names of Afrotropical species: Billaea rubida O’Hara & Cerretti for Phorostoma rutilans Villeneuve, 1916, preoccupied in the genus Billaea Robineau-Desvoidy, 1830 by Musca rutilans Fabricius, 1781, nom. n.; Cylindromyia braueri O’Hara & Cerretti for Ocyptera nigra Villeneuve, 1918, preoccupied in the genus Cylindromyia Meigen, 1803 by Glossidionophora nigra Bigot, 1885, nom. n.; Cylindromyia rufohumera O’Hara & Cerretti for Ocyptera scapularis Villeneuve, 1944, preoccupied in the genus Cylindromyia Meigen, 1803 by Ocyptera scapularis Loew, 1845, nom. n.; Phytomyptera longiarista O’Hara & Cerretti for Phytomyzoneura aristalis Villeneuve, 1936, preoccupied in the genus Phytomyptera Rondani, 1845 by Phasiostoma aristalis Townsend, 1915, nom. n.; and Siphona (Siphona) pretoriana O’Hara & Cerretti for Siphona laticornis Curran, 1941, preoccupied in the genus Siphona Meigen, 1803 by Actia laticornis Malloch, 1930, nom. n. New type species fixations are made under the provisions of Article 70.3.2 of the ICZN Code for two genus-group names: Lydellina Villeneuve, 1916, type species newly fixed as Lydellina villeneuvei Townsend, 1933 (valid genus name); and Sericophoromyia Austen, 1909, type species newly fixed as Tachina quadrata Wiedemann, 1830 (synonym of Winthemia Robineau-Desvoidy, 1830). Lectotypes are designated for the following nine nominal species based on examination of one or more syntypes of each: Degeeria crocea Villeneuve, 1950; Degeeria semirufa Villeneuve, 1950; Erycia brunnescens Villeneuve, 1934; Exorista oculata Villeneuve, 1910; Kiniatilla tricincta Villeneuve, 1938; Myxarchiclops caffer Villeneuve, 1916; Ocyptera linearis Villeneuve, 1936; Peristasisea luteola Villeneuve, 1934; and Phorocera crassipalpis Villeneuve, 1938. The following four genus-group names that were previously treated as junior synonyms or subgenera are recognized as valid generic names: Bogosiella Villeneuve, 1923, status revived; Dyshypostena Villeneuve, 1939, status revived; Perlucidina Mesnil, 1952, status revived; and Thelymyiops Mesnil, 1950, status n. The following six species-group names that were previously treated as junior synonyms are recognized as valid species names: Besseria fossulata Bezzi, 1908, status revived; Degeeria cinctella Villeneuve, 1950, status revived (as Medina cinctella (Villeneuve)); Nemoraea miranda intacta Villeneuve, 1916, status revived (as Nemoraea intacta Villeneuve); Succingulum exiguum Villeneuve, 1935, status revived (as Trigonospila exigua (Villeneuve)); Wagneria rufitibia abbreviata Mesnil, 1950, status n. (as Periscepsia abbreviata (Mesnil)); and Wagneria rufitibia nudinerva Mesnil, 1950, status n. (as Periscepsia nudinerva (Mesnil)). The following 25 new or revived combinations are proposed: Afrophylax aureiventris (Villeneuve, 1910), comb. n.; Blepharella orbitalis (Curran, 1927), comb. n.; Bogosiella pomeroyi Villeneuve, 1923, comb. revived; Brachychaetoides violacea (Curran, 1927), comb. n.; Carceliathrix crassipalpis (Villeneuve, 1938), comb. n.; Charitella whitmorei (Cerretti, 2012), comb. n.; Dyshypostena edwardsi (van Emden, 1960), comb. n.; Dyshypostena tarsalis Villeneuve, 1939, comb. revived; Estheria buccata (van Emden, 1947), comb. n.; Estheria surda (Curran, 1933), comb. n.; Filistea aureofasciata (Curran, 1927), comb. n.; Madremyia setinervis (Mesnil, 1968), comb. n.; Mesnilotrix empiformis (Mesnil, 1976), comb. n.; Myxophryxe longirostris (Villeneuve, 1938), comb. n.; Nealsomyia chloronitens (Mesnil, 1977), comb. n.; Nealsomyia clausa (Curran, 1940), comb. n.; Nilea longicauda (Mesnil, 1970), comb. n.; Paratrixa aethiopica Mesnil, 1952, comb. revived; Paratrixa stammeri Mesnil, 1952, comb. revived; Perlucidina africana (Jaennicke, 1867), comb. n.; Perlucidina perlucida (Karsch, 1886), comb. revived; Prolophosia retroflexa (Villeneuve, 1944), comb. n.; Sturmia profana (Karsch, 1888), comb. n.; additionally, Ceromasia rufiventris Curran, 1927 is treated as an unplaced species of Goniini, comb. n. and Hemiwinthemia stuckenbergi Verbeke, 1973 is treated as an unplaced species of Leskiini, comb. n. New or revived generic and specific synonymies are proposed for the following nine names: Afrosturmia Curran, 1927 with Blepharella Macquart, 1851, syn. n.; Archiphania van Emden, 1945 with Catharosia Rondani, 1868, syn. revived; Besseria longicornis Zeegers, 2007 with Besseria fossulata Bezzi, 1908 (current name Besseria fossulata), syn. n.; Dexiomera Curran, 1933 with Estheria Robineau-Desvoidy, 1830, syn. n.; Hemiwinthemia francoisi Verbeke, 1973 with Nemoraea capensis Schiner, 1868 (current name Smidtia capensis), syn. n.; Kinangopana van Emden, 1960 with Dyshypostena Villeneuve, 1939, syn. n.; Metadrinomyia Shima, 1980 with Charitella Mesnil, 1957, syn. n.; Phorocera majestica Curran, 1940 with Phorocera longirostris Villeneuve, 1938 (current name Myxophryxe longirostris), syn. n.; and Podomyia discalis Curran, 1939 with Antistasea fimbriata Bischof, 1904 (current name Antistasea fimbriata), syn. n.

Review of Parasitoid Wasps and Flies (Hymenoptera, Diptera) Associated with Limacodidae (Lepidoptera) in North America, with a Key to Genera

Abstract. Hymenopteran and dipteran parasitoids of slug moth caterpillars (Lepidoptera: Limacodidae) from North America are reviewed, and an illustrated key to 23 genera is presented. Limacodid surveys and rearing were conducted during the summer months of 2004–2009 as part of research on the ecology and natural history of Limacodidae in the mid-Atlantic region of the U.S.A. Parasitoid rearing involved a combination of collecting naturally occurring larvae in the field (at least 14 host species) and placing out large numbers of “sentinel” larvae derived from laboratory colonies of three host species. Species in the following families are documented from limacodids in North America as primary or secondary parasitoids (number of genera for each family in parentheses; number of genera included in key but not reared through this research in brackets): Chalcididae ([1]; Hymenoptera: Chalcidoidea), Eulophidae (3; Chalcidoidea), Pteromalidae ([1]; Chalcidoidea), Trichogrammatidae (1; Chalcidoidea), Braconidae (3 [1]; Hymenoptera: Ichneumonoidea), Ichneumonidae (7 [3]; Ichneumonoidea), Ceraphronidae (1; Hymenoptera: Ceraphronoidea), Trigonalidae (2; Hymenoptera: Trigonaloidea), Bombyliidae ([1]; Diptera: Asilioidea), and Tachinidae (3; Oestroidea). We recovered 20 of 28 genera known to attack limacodids in North America. Records discerned through rearing in the mid-Atlantic region are augmented with previously published host-parasitoid relationships for Limacodidae in North America north of Mexico. New records are reported for the following parasitoids (total new records in parentheses): Uramya limacodis (Walker) (1), U. pristis (Townsend) (5), Austrophorocera spp. (6), Ceraphron sp. (1), Alveoplectrus lilli Gates (1), Playplectrus americana (Girault) (10), Pediobius crassicornis (Thomson) (1), Trichogramma (1), Mesochorus discitergus (Say) (1), Hyposoter fugitivus (Say) (1), and Isdromas lycaenae (Howard) (5). The male of Platyplectrus americana (Hymenoptera: Eulophidae) is redescribed, and the female is described for the first time. Incidental and miscellaneous host-parasitoid associations are discussed, and it is concluded that most of these records are likely parasitoids of contaminants accidentally introduced during the limacodid rearing process. Triraphis eupoeyiae (Ashmead), new combination, is transferred from Rogas (Hymenoptera: Braconidae).

Survey of first instars of the Siphonini (Diptera: Tachinidae)

First instars of 68 species of the Siphonini were examined, including representatives of all genera of the tribe except Proceromyia Mesnil, and all subgenera of Siphona Meigen except Uruactia Townsend. Two aspects of first instar siphonines are markedly varied: features of the cephalopharyngeal skeleton and vestiture ventrally on posterior abdominal segments. These regions of first instars are described for the examined genera and subgenera of the Siphonini, in addition to four groups of undescribed, New World species of Siphona s.l. unplaced to subgenus. The phylogenetic significance of particular character states of first instar siphonines is discussed.

Correlation between wing size and position of a hind crossvein in the Siphonini (Diptera: Tachinidae)

In the Siphonini (Diptera: Tachinidae), crossvein dm-cu is relatively near the wing margin (the primitive condition within the tribe) in larger siphonines and displaced toward the middle of the wing (derived condition) in smaller siphonines. Displacement of dm-cu increases the passively deformable area of the wing relative to the supporting area, presumably increasing the aerodynamic efficiency of flight in smaller siphonines. The trend toward increasing the passively deformable area in smaller siphonines compared with larger ones is complementary to the general trend toward reduced venation in tiny insects.

The History of Dipterology at the Canadian National Collection of Insects, with Special Reference to the Manual of Nearctic Diptera

Abstract The history of Diptera research at the Canadian National Collection of Insects is briefly outlined. Short biographic sketches of the coordinators of the Manual of Nearctic Diptera are given and the development of the Manual project is presented to provide background on their achievements. Lists of publications by each of the coordinators and of patronyms honouring them are provided. This Festschrift honours the remarkable contributions of the coordinators, J. Frank McAlpine, Bobbie V. Peterson, Guy E. Shewell, Herbert J. Teskey, J. Richard Vockeroth, and D. Monty Wood.

Molecular phylogenetics and piercer evolution in the bug-killing flies (Diptera: Tachinidae: Phasiinae): Phasiine phylogenetics and piercer evolution

Phasiinae (Diptera: Tachinidae) are endoparasitoid flies that attack Heteroptera, including a multitude of agricultural pests. A phylogenetically informed classification of Phasiinae has eluded systematists for over a century, primarily because of the conflicting character states and confusing morphology of certain taxa that indicate potential placement within other subfamilies. The unstable nature of phasiine taxonomy discourages important research into their classification, life history and potential use in biological control. In hopes of resolving several longstanding taxonomic debates and encouraging future research into this important group of parasitoids, the first molecular systematic analysis of Phasiinae is presented, including 128 worldwide taxa (80 genera) and approximately 7.6 kb of nuclear data representing four genes. Special emphasis is placed on the resolution of taxonomically ambiguous groups. The resulting robustly supported phylogenetic trees [maximum‐likelihood (ML)/Bayesian] were used to trace the evolution of significant adaptive traits within Tachinidae and test hypotheses about the classification of Phasiinae. Subfamily placements of certain taxa are confidently resolved including Eutherini, Epigrimyiini, Litophasia Girschner within Dexiinae, and Strongygastrini and Parerigonini within Phasiinae. The members of tribe Phasiini are redistributed: Cistogaster Latreille, Clytiomya Rondani, Ectophasia Townsend, Eliozeta Rondani and Euclytia Townsend transferred to Gymnosomatini; Opesia Robineau‐Desvoidy to Strongygastrini; and Xysta Meigen to Xystini. Similarly, members of Parerigonini are treated as belonging to Parerigonini (Parerigone Brauer, Zambesomima Walker), Cylindromyiini (Australotachina Curran, Pygidimyia Crosskey, Neobrachelia Townsend) or new tribe Zitini (Zita Curran, Leverella Baranov). Penthosia van der Wulp is transferred from Cylindromyiini to Hermyini. Ancestral state reconstruction suggests that piercing structures used to insert eggs directly into host tissues have evolved separately in a number of groups, but have also been lost or reduced in several lineages. A single potentially unequivocal morphological synapomorphy of Phasiinae, an elongated medial plate of the hypandrium in males, is identified.

The identity of Tachina westermanni Wiedemann, 1819 (Diptera: Calliphoridae or Tachinidae) with a solution to a nomenclatural problem

Tachina westermanni Wiedemann, 1819 was based on four syntypes, two conspecific calliphorids and two conspecific tachinids. Two existing but contradictory lectotype fixations have resulted in confusion as to the correct application of the specific name westermanni Wiedemann. Evidence is presented showing that the lectotype fixation of Townsend in 1931 is valid and assigns westermanni Wiedemann to the Calliphoridae, with Pericallimyia westermanni as the valid binomen. The valid name for the tachinid taxon becomes Brachelia westermanni Robineau-Desvoidy, 1830 and a neotype is designated for it in the interests of nomenclatural stability.